Macroinvertebrate Colonization Dynamics on Artificial Substrates
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Hydrobiologia (2014) 727:1–18 DOI 10.1007/s10750-013-1779-z PRIMARY RESEARCH PAPER Macroinvertebrate colonization dynamics on artificial substrates along an algal resource gradient A. Braccia • S. L. Eggert • N. King Received: 13 August 2013 / Revised: 1 December 2013 / Accepted: 2 December 2013 / Published online: 14 December 2013 Ó Springer Science+Business Media Dordrecht 2013 Abstract Riparian canopy removal and land use also increased according to the gradient. Correlation may introduce multiple stressors that can alter food results indicated that chlorophyll a biomass, rather and habitat for stream organisms, but the influence of than time or temperature, was better related to the these alterations on macroinvertebrate colonization abundance and biomass of most primary consumers on dynamics is less well known. A field study involving substrates. These results suggest that the combined the simultaneous placement and removal of artificial effects of elevated temperatures and nutrients can substrates was performed to examine how macroin- result in relatively rapid algal accrual that may alter vertebrate colonization rates might vary with algal the colonization and establishment of macroinverte- accumulation within a perennial stream segment in brate communities in streams subjected to gradients of eastern Ohio, USA. The study was conducted over a riparian disturbances. 60-day summer colonization period in three reaches that were selected to represent an algal resource Keywords Functional feeding groups Á gradient according to canopy cover and agricultural Chlorophyll a Á Periphyton Á Nutrients Á nutrient sources in the riparian corridor. Total nitro- Agriculture Á Riparian canopy gen, water temperatures, and mean algal biomass from substrates increased along the resource gradient rep- resented by the study sites. Total macroinvertebrate Introduction biomass and the abundance and biomass of scrapers Identifying factors that influence the colonization dynamics of stream macroinvertebrates can improve Handling editor: Sonja Stendera our understanding of stream recovery following A. Braccia (&) natural and anthropogenic disturbance and help Department of Biological Sciences, Eastern Kentucky explain temporal variability in community structure University, Richmond, KY 40475, USA that is commonly observed in benthic macroinverte- e-mail: [email protected] brate studies. Since the degree and extent of distur- S. L. Eggert bance determines the size of habitat patches available USDA Forest Service, Northern Research Station, Grand for colonization, studies of macroinvertebrate coloni- Rapids, MN 55744, USA zation are conducted over large (e.g., rivers, streams; Williams & Hynes, 1977; Gray & Fisher, 1981; N. King Biology Department, Bucknell University, Lewisburg, Minshall et al., 1983a; Malmqvist et al., 1991; Milner PA 17837, USA et al., 2000) and small (e.g., stones, patches; Wise & 123 2 Hydrobiologia (2014) 727:1–18 Molles, 1979; Shaw & Minshall, 1980; Peckarsky, 1980; Williams, 1980; Robinson et al., 1993; Maton- 1986; Robinson et al., 1990) spatial scales. The ickin et al., 2001). Simultaneous measures of basal simultaneous placement and removal of artificial food resources (e.g., algae and detritus) have not substrates is an approach that has been widely used always been reported from colonization studies, so it is to study macroinvertebrate colonization of benthic difficult to generalize about how the accumulation of substrate patches following small-scale disturbance, these resources influence macroinvertebrate commu- such as flow-generated scour or tumbling of rocks nity colonization. However, continuous increases in (Ciborowski & Clifford, 1984; Lake & Doeg, 1985; macroinvertebrate densities and richness have been Robinson & Minshall, 1986; Downes & Lake, 1991; attributed to changes in habitat and the accumulation Rutherford, 1995; Baer et al., 2001; Miyake et al., of food resources on substrates over colonization 2003). intervals (Meier et al., 1979; Shaw & Minshall, 1980; The succession of taxa and trophic groups onto Baer et al., 2001), and more rapid colonization of benthic habitats has been well described from macr- substrates by macroinvertebrates has been reported oinvertebrate colonization studies. In general, highly from studies that used substrates with natural or mobile, opportunistic taxa within the collector-filter- artificial periphyton layers (Robinson et al., 1990; ing or -gathering functional groups will colonize first Baer et al., 2001; Miyake et al., 2003). Further, and then increases in the abundance of scrapers, Robinson et al. (1990) suggested that variable stabil- predators, and shredders can occur (see review by ization times for macroinvertebrate densities from Mackay, 1992). Colonization models predict that early small-scale colonization studies may be partly dynamics are primarily a result of stochastic factors explained by the presence of a periphyton layer. Since and that initial colonization should quickly increase to algal resources may provide food and habitat for maximum abundance, biomass, or taxa richness, and certain macroinvertebrates, equilibrium is unlikely to then remain at an equilibrium level (Sheldon, 1984; develop if substrates are continuously accumulating Minshall & Petersen, 1985). Time to equilibrium (i.e., epilithic material (Mackay, 1992). stabilization time) is often equated with plateaus (or Benthic food resources can be altered in stream maxima) in macroinvertebrate densities, biomass, or reaches where native forest has been removed from taxa richness. Results from small-scale colonization riparian zones because increased solar radiation and studies have shown that initial colonization is rapid temperatures can enhance standing stocks of periph- (2–4 days), but individuals of different taxa and yton and primary production (Lowe et al., 1986; Hill trophic groups colonize at different rates, and time to et al., 1995; Quinn et al., 1997; Kiffney et al., 2003). equilibrium in densities and richness is variable and Riparian canopy removal also reduces forest litter generally ranges from 4 to 30 days (Rosenberg & inputs that support the detrital-based food webs of Resh, 1982; Mackay, 1992). In some studies, equilib- temperate streams, and results from field studies have rium in total macroinvertebrate densities or richness shown that timber harvest from riparian zones can on substrates was never reached and successional increase the abundance of macroinvertebrates and changes in individual taxa extended beyond stabiliza- alter the functional composition of macroinvertebrate tion times for total densities or richness (Meier et al., communities (Newbold et al., 1980; Hawkins et al., 1979; Peckarsky, 1986; Baer et al., 2001; Miyake 1982; Wallace & Gurtz, 1986). In addition, agricul- et al., 2003). tural land use within watersheds can introduce inor- Variation in stabilization times for macroinverte- ganic nutrients (Broussard & Turner, 2009; Hill et al., brate densities has been attributed to season, experi- 2011) that may interact with light and elevated mental design, and the densities, mobility, and life temperatures and further stimulate primary produc- histories of available colonists (Rosenberg & Resh, tion. Altered riparian zones and nutrient inputs have 1982; Robinson et al., 1993; Miyake et al., 2003). been identified as major stressors to benthic macroin- Studies from temperate streams have demonstrated vertebrate communities (Dodds & Welch, 2000; that colonization rates of macroinvertebrates are more Paulsen et al., 2008). Results from the analysis of rapid during summer and early fall when elevated large observational datasets have revealed that macr- stream temperatures increase macroinvertebrate activ- oinvertebrate-based indicators of biological integrity ity and more recruits are present (Shaw & Minshall, (e.g., EPT taxa, total taxa richness) are negatively 123 Hydrobiologia (2014) 727:1–18 3 correlated with nutrients in wadeable streams in some region receives an average of 96 cm of precipitation regions of the U.S. (Miltner & Rankin, 1998; Wang per year. Based on coverage from the 2006 National et al., 2007; Yuan, 2010). Understanding the relative Land Cover Database (Homer et al., 2007), the role of algal resource accrual on macroinvertebrate watershed is predominantly forested (49%) with colonization dynamics may reveal mechanisms that cultivated crops and pasture (41%) and low/medium influence the structure of benthic macroinvertebrate intensity development (9%). Land use in the riparian communities in agricultural streams. We are unaware corridor is variable and areas of agricultural and light of any small-scale colonization studies that have residential use are interspersed among patches of addressed the question of how macroinvertebrate continuous forest. colonization of artificial substrates might be influ- The ‘‘low’’ end of the expected resource gradient enced by the combined effects of reduced riparian has a forested riparian zone (Forested Site) and is canopy cover and agricultural land use. located at the Hiram College’s James H. Barrow Field Since primary productivity can be enhanced by Station. The field station contains 200 acres of beech- light, temperature and nutrients, and macroinverte- maple forest and no agricultural nutrient sources. In brate colonization dynamics can be influenced by algal 2006, the Forested Site was classified as Coldwater accrual on substrates, we hypothesized that macroin- Habitat for fish and macroinvertebrates and fully vertebrate